New takes on the Wealden Supergroup palaeobiota, part 2: Baryonyx, freshwater plesiosaurs, ornithomimosaurs and others

Last week we took a look at some new art of animals from the Wealden Supergroup, the intensively studied, historically important Lower Cretaceous rocks of Southern Britain. We all know the Wealden for celebrity dinosaurs like Iguanodon and Baryonyx, but there's a heap of other interesting animals in there which get relatively little publicity. It's mostly these we're focusing on here, in the second (and final) part of these 'picture of the day'-style posts. 

As before, if you like anything here, remember that you can buy prints of them all from my shop (the Wealden section might be relevant) and its new Facebook outlet. Indeed, if you like my work and are on Facebook, why not 'like' the new Mark Witton Palaeoart page? It's the best place to see when new prints and finished pictures are available.

Baryonyx walkeri: king of the fishers, redux

Baryonyx walkeri, off for a stroll among the crocodyliforms and pterosaurs.

Let's break this post in with a familiar animal: spinosaurid Baryonyx. It's hard to appreciate now how weird this animal seemed back in the 1980s and 1990s. At this point, other spinosaur material was only very poorly known, and laymen and scientists alike found this weird, superficially-crocodile like animal fascinating. Ironically, it's recently turned out that we first collected Wealden spinosaur material centuries ago, but struggled to recognise its significance until more complete remains were unearthed in the 1980s. We now know that Baryonyx can be found throughout a good chunk of upper Wealden stratigraphy and teeth referable to it - or another spinosaurid - are fairly common, at least as Wealden dinosaur fossils go. Baryonyx provided the basic template we'd recognise for all spinosaurid anatomy until last year when, famously, some spinosaurs were proposed to be rather different. It's clear that, whatever is going on with Spinosaurus, Baryonyx retains more conventional hindlimb and pelvic proportions, and may not have been so aquatically adapted as true spinosaurines. In this updated image, B. walkeri is splashing into a body of water while goniopholidid crocodyliforms and gnathosaurine pterosaurs go about their business around it. Note how much larger Baryonyx is compared to the crocs: Baryonyx is the largest theropod in the Wealden Supergroup, by a good margin.

Button-toothed crocs, redux

Bernissartid Koumpiodontosuchus aprosdokiti foraging for molluscs. It's eating a mud snail, Viviparus cariniferus, while tiny (6 mm long) physid gastropods Prophysa crawl over pond scum in the lower left of the image. Dragonflies provide scale, and unnamed tetanurans prowl around the background.

Last year I was lucky enough to provide the first restoration of Kompiodontosuchus aprosdokiti, a small neosuchian crocodyliform common to the Wessex Formation, and perhaps other parts of the Wealden sequence. Koumpiodontosuchus is a bernissartid, a group of small-bodied crocodyliforms with robust, shell-cracking teeth at the back of their jaws. As you'll know if you read my write up last year, these were likely employed in smashing molluscs and insects. The tetanuran theropods in this image are unnamed, but are not thought to be referable to any existing Wealden taxa. We probably need more material of them to consider them nameable, however: recognising that they are different from other Wealden theropods is only half the battle. Modern students of Wealden fossils famously do their best to preserve historic names based on fragmentary bones, but there seems to be an effort to 'future proof' Wealden taxonomy against confusion by only naming well-represented, characteristic animals. I guess I could have chosen one of the better known theropods to play the 'This was the Age of Dinosaurs' card for this PR image, but I think it's good to show that not all large theropods in the Wessex palaeobiota were Neovenator, Baryonyx or Eotyrannus. 

Welcoming the new Wealden ornithomimosaurs

A flock of Wessex Formation ornithomimosaurs forage in a marshland, while istiodactylid pterosaurs skulk about behind them.

Those keeping their ears to the ground will know that the newest arrivals to the Wealden dinosaur palaeobiota are ornithomimosaurs, commonly known as ostrich dinosaurs. Two specimens show that these animals were present in both the Weald and Wessex basins of the broader Wealden succession, and one of these fossils represents a historic taxon named in 1889: Valdoraptor oweni. Key to identifying ostrich dinosaurs in the Wealden was the discovery of abundant ornithomimosaur remains in France, many of which are so reminiscent of Valdoraptor and other Wealden theropod material that they may represent the same taxon. If you want to know more about these and their relationship to the complex story of Wealden theropods, check out Darren Naish's post on this at Tetrapod Zoology.

The above new painting shows a group of (nameless) Wessex Formation ornithomimosaurs in a well-vegetated marshland, in the rainy season, while istiodactylid pterosaurs mosey about in the background. The abundance of ostrich dinosaurs and juveniles in the middle-right are nods to the frequent recovery of abundant specimens of different levels of maturity at many ostrich dinosaur sites, including the new, French 'Angeac ornithomimosaur'. Note that the wings of the running foreround animal are somewhat swept back: I don't think the more common way of reconstructing ornithomimosaurs with 'dangly arms' looks right. They look like they should be holding shopping bags or something.

Valdosaurus in the forest, redux

Two Wealden dryosaurids Valdosaurus canaliculatus, and a stubborn avialan.

Ornithomimosaurs weren't the only fast runners in Wealden landscapes. Dryosaurids, like Valdosaurus canaliculatus were also fleet-footed animals with powerful, well-muscled hindlimbs, and tiny bodies attached to the front. In this reworked image, two of these 3-4 m long animals are taking it slow through a Wealden woodland. Although Wealden climates were quite warm and arid, leaving much of the landscape looking quite chaparral-like, some relatively upland parts seem to have been more vegetated: it's here that this picture is set. In my mind, these animals always walked with the stooping posture of the foreground animal - as noted last time, I like the idea that prehistoric animals had characteristic postures varying slightly from those we consistently restore in skeletal restorations. Note the avialan on the left of the image, which is a nod to the recovery of bird teeth from Wealden deposits. Anyone who's ever been forced to walk around a stubborn reclined mallard will recognise the situation now facing the Valdosaurus.

Barilium dawsomi in leathers, redux

Barilium dawsoni, a large and very robust iguanodont from Sussex. A flock of 'Ashdown maniraptorans' add scale.

Last time we featured Iguanodon bernissartensis: now it's the turn of the 'other' big Wealden iguanodont, the stratigraphically older, and osteologically chunkier Barilium dawsoni. In this redone painting, I've tried to make the Barilium skin more interesting than just plain old scales, covering the back in small, horny ossicles and creasing the flanks as if the skin is particularly thick, leathery and folded. I think we should be rendering more interesting skin regularly in scaly dinosaur palaeoart, as it seems most extensive dinosaur skin remains show unexpected features - strangle scales, wattles, folds and that sort of thing - which small skin patches mostly cannot record adequately. It's interesting to contrast these skin impressions with homogeneous restorations of scaly dinosaur appearance presented by some, where every species is covered in smooth hide following perfect contours of the underlying tissues: I'm not sure that's what fossils are telling us. As before, the 'Ashdown maniraptoran' provides scale to the bulk of Barilium. For the uninitiated, the Ashdown maniraptoran is seriously small for a Mesozoic dinosaur - maybe about 30-50 cm long. If you find big iguanodonts exciting, be sure to check out this previous post.

Polacanthus redux, again

A Wealden tree vies for attention with Polacanthus foxii, and some tiny birds.

OK, I'm cheating a bit with this one. This redone version of a much older painting has been posted fairly recently, but it seemed a bit remiss to skip this ankylosaur in this run down of recently produced Wealden palaeoart. Polacanthus foxii is, of course, the Wealden's sacral-shield-bearing nodosaurid, shown here strolling around a Cretaceous hillock with some birds for company. Having scratched the completist itch, let's move on, because we've seen this all before.

Accidentally sinister Leptocleidus, redux 

Mother and calf Leptocleidus superstes, a freshwater leptocleidid plesiosaur, explore a river inlet in Lower Cretaceous Sussex. 

Our final stop is in Wealden rivers and estuaries, where Leptocleidus superstes and other species of freshwater leptocleidid plesiosaurs roamed. The new version of this image has added a lot of detail on top of the original, which has inadvertently made the mother and calf Leptocleidus look more sinister than intended - hey, it's not my fault their teeth stick out like that. Back in the original post on these animals I mentioned that pliosaurs may also have been present in Wealden lakes and rivers, but note that this is no longer certain: the Hastanectes valdensis remains once provisionally considered pliosauroid have been placed in Leptocleididae in more recent analyses. That does make for a neater story - it means that leptocleidids retain their dominant role as 'near-shore-or-freshwater' animals, but perhaps a slightly less interesting one.

And that's all for now - I hope you've enjoyed this jaunt back to the ancient Wealden and these revised artworks. I'm sure we'll visit the Wealden again in time. Coming next, probably: walking with non-pterodactyloid pterosaurs.

New takes on the Wealden Supergroup palaeobiota, part 1: Iguanodon, Neovenator, Eotyrannus and others

Regular readers will know that I'm prone to dabbling in palaeoart depicting the environments and animals of the Wealden Supergroup, the 18 million year stretch of Early Cretaceous time represented by mud-and sandstone deposits across the southern UK. Recently, I've been updating some existing Wealden work as well as producing some new stuff of other Wealden species. With no time to produce a new post of substance, here's a bumper 'picture of the day'-type post. Initially, I was going to chuck something like ten images on here, but time has run short and I'll have to split it in two.

If you like anything here, remember that you can buy prints of them all from my shop (there's now a Wealden section, too), which is now also browsable from the comfort of Facebook. OK, enough preamble: into the Wealden once again...

Iguanodon bernissartensis: thumb wars

Two Iguanodon bernissartensis, the quintessential Wealden iguanodont, decide to settle their differences, while members of their herd watch on.

Poor old Iguanodon doesn't get the attention it used to, and a lot palaeoart we do see of it tends to focus on tried and tested behaviours: lots of standing about and eating, but not much else. In this new painting, I've attempted to show two big Iguanodon individuals settling an intra-specific dispute via use of thumb spikes. Long-term readers may recall that we've covered iguanodont thumb spikes before, and that I. bernissartensis has especially big ones. Here, they've been swinging their thumbs at each other's soft bits, causing deep, bloody wounds. This might seem extreme, but there are plenty of modern animals which take intraspecific fights to similarly gory levels - elephant seals were a key inspiration here. I imagine battling Iguanodon would look like an armed sumo-wrestling match, albeit with longer tails and less rice. Note that you can see the breath of several animals here: Wealden winters are not meant to be especially warm.

Rebbachisaurids vs. Neovenator salerii redux

Carcharodontosaurian Neovenator salerii stalks a pair of rebbachisaurid sauropods, using darkness as cover.

A while back I posted about dinosaur predation, noting that modern animal predator acts are often far less gladiatorial and epic than we might imagine. It's this slow, considered approach to predation which I'm attempting to show here, as the carcharodontosaur Neovenator stalks two rebbachisaurid sauropods in the dead of night. The idea is that the Neovenator has much better eyesight than the sauropods, who know they're in trouble, but can't really respond adequately. Note the rain: some recent models of Wealden palaeoclimates suggest it was wetter than previously modelled (albeit with very high evaporation rates for much of the year).

Anteophthalmosuchus hooleyi vs. Hypsilophodon foxii, redux

Large goniopholidid Anteophthalmosuchus hooleyi takes advantage of a flooding river to hunt two stranded Hypsilophodon foxii.

Speaking of rain, we know that some parts of the Wealden were prone to flooding following particularly intense downpours. That's good news for animals adapted for powerful swimming, but less welcome to species which prefer dry land. Here, in this reworked painting, the large Wealden goniopholidid Anteophthalmosuchus hooleyi has found a stranded pair of adult and juvenile Hypsilophodon foxii, and is taking full advantage of the situation. Goniopholodids are a group of almost-crocodiles characterised by long forelimbs, interlocking scutes and overbitten jaws - you can read more about them here.

Eotyrannus lengi: firestarter, redux 

Early tyrannosauroid Eotyrannus lengi stalks the edge of such a wildfire. 

What else does rain bring? Sometimes, lightning. When introduced to a parched Wealden landscape, lightning strikes caused short-lived canopy fires which, ultimately, created conditions ideal for fossil preservation. In this reworked painting, a fully-feathered tyrannosauroid Eotyrannus lengi is prowling the periphery of a Wealden canopy fire to grab any animals flushed out by the flames.

The tiny wars of Wesserpeton evansae, redux 

Two Wesserpeton evansae get in each other's faces, because some animals are just jerks.

OK, enough about Wealden weather. Here's a reworked version of two of the Wealden's tiniest tetrapods - indeed, some of the smallest fossil tetrapods of all - facing off in leaf litter. Recently named Wesserpeton evansae, these are albanerpetontids, very small amphibians which only died out a few million years ago. The 35 mm snout-vent length of these animals did nothing to temper their ferocity, and numerous jaws of Wesserpeton have healed fractures and breaks from intraspecific tussles. The animals in this picture are speaking the aggressive body language of modern salamanders as a prelude to their conflict. Two sauropods hang around in the background because, hey, it's called the Age of Dinosaurs for a reason. Some people have suggested this image borders on the trippy and surreal. Stay off the shrooms, kids. 

Rebbachisaurids and chums

Lower Cretaceous rebbachisaurids and giant sauropod 'Angloposeidon' look for water in this desiccating Wealden lake.

I do like rebbachisaurids, that group of sauropods who didn't get the memo about long necks. They're only represented by scrappy remains in the Wealden (a scapula) which is enough to tell us they were there, but not substantial enough to carry a name. Here, a few individuals are digging around a rapidly drying lake-bed to find a substantial source of water: digging elephants were the inspiration for this scene. In the background, probable brachiosaurid 'Angloposeidon' struts its stuff. It's meant to be walking particularly tall - I like the idea that fossil animals would carry themselves in different, characteristic ways, just as modern animals do. A pink gnathosaurine pterosaur has snuck into the foreground, just because. 

A lesser-seen Wealden scene: the Hastings Beds palaeobiota

Finally for now, here's one more new painting. This is a reconstruction of a swollen river representing part of the Hastings Beds, the oldest deposits of the Wealden, complete with local reptile fauna. The animals shown here are really poorly known: titanosaur 'Pelorosaurus' becklesii (bits of forelimb), possible carcharodontosaurian Becklespinax altispinax (three dorsal vertebrae), eucryptodiran turtle Hylaeochelys belli (a shell), and the possible azhdarchoid previously known as 'Palaeornis cliftii' (humerus). So yes, take the 'restorations' of these animals with an evaporite mine of salt: they're really just better known, fairly 'generic' representatives of groups represented by these Wealden taxa, air-dropped into a Wealden setting. Becklespinax is obviously modelled closely on Concavenator, as they seem to be pretty closely related and have a similar taste in dorsal ornamentation. I gave Becklespinax a more vertical anterior sail margin however, as indicated by the fossil. There's an article waiting to be written on palaeoart like this - should we even bother 'reconstructing' poorly known scenes and species? I clearly think we should, but we'll have to discuss the reasons why another time. 

I'm just now realising that there's a lot of confrontation in these images. Come back soon for a more placid, relaxed set of pictures in part 2...

Remembering Iguanodon

Retrosaur Iguanodon, c. 1854. Based, of course, on the sublime work of Benjamin Waterhouse Hawkins.

Space year 2014 marks the 189th anniversary of the naming of a dinosaur icon, Iguanodon. The major beats of the discovery and research history of this Lower Cretaceous herbivore are so well-established within palaeontological lore that most readers will need little reminder of it here. We all know that Iguanodon was first known from large, iguana-like teeth found in southern England in 1822, supposedly by Gideon Mantell's wife, which we all also know is widely considered an embellished tale: the teeth were probably found by Mantell himself or quarrymen. Equally familiar is Mantell's naming of Iguanodon in 1825 with the first specific name given to this genus, anglicus, added by Friedrich Holl in 1829. As the second dinosaur to be named, Iguanodon was part of the trio of dinosaur genera used by Richard Owen to name Dinosauria in 1842 and was reconstructed alongside its cousins, Megalosaurus and Hylaeosaurus, by Richard Owen and Benjamin Waterhouse Hawkins as an awesome dinosaurian rhino in 1854. Discoveries of more complete Iguanodon remains, first in Britain and then in the coal mines of Bernissart, Belgium, led to a reconsideration of this bauplan. The most extensive work on this front was conducted by Louis Dollo in the 1880s, who took the complete Iguanodon skeletons from Bernissart - among the first complete dinosaurs known from anywhere in the world at that time - and created the famous 'kangaroo' posture for Iguanodon, broken tails and all, which dominated reconstructions of this animal for the next century. It was not until the 1980s that Iguanodon adopted the appearance of the facultatively bipedal, horizontally-backed ornithopod we know today. So far, so familiar.

Undoubtedly, Iguanodon is a 'classic' dinosaur, and has been a mainstay of popular dinosaur literature for decades. Other dinosaur species named in the early 1800s have not enjoyed the same treatment (Thecodontosaurus, Ceitiosaurus and Hylaeosaurus for instance, are not household names), so its popularity is not just a result of it being one of the first dinosaurs known. Most of us can probably remember a key Iguanodon depiction from our childhood dinosaur books, magazines or films - or from a Love in the Time of Chasmosaurs vintage palaeoart post if you're not yet through puberty - with it stood upright and, of course, giving an irrepressible thumbs-up with its famous thumb spike. These Mesozoic Fonzies, which diehards always knew came in big (I. bernissartensis) and small (I. atherfieldensis) flavours, wouldn't stop manually approving their surroundings even when being attacked by passing generic 'megalosaurs'. Final revisions to its anatomy - an aloft tail and quadrupedal stance - have been drifting into popular depictions for years now, replacing MesoFonz with a heavyset herbivore often depicted living in herds and browsing at different levels. While its lack or truly bizarre anatomy or ferocity may have prevented Iguanodon from ever being the most famous of dinosaur species, there's little doubt that it's held a long-term place in palaeo-pop culture.

All good things...
At least, until recently. If the internet palaeo scene is anything to go by, Iguanodon seems to be sliding down the popularity pole at the moment. It just doesn't seem to be the topic of much conversation any more, or even artwork. Feathered theropods, weird sauropods, horned dinosaurs and even hadrosaurs - boring old hadrosaurs - seem to have stolen the limelight. Perhaps this is because our taxonomic and palaeobiological perceptions of many prehistoric animals have radically changed in recent years whereas Iguanodon, frankly, has remained rather static. It's a bit too familiar. Dinosaur palaeontology has changed radically in the last few decades, but it's changed around Iguanodon, which has done little more than tip forward a little since the 1980s. Discussions about feathers, postures, weird soft-tissue details and whatnot have passed it by entirely, and even a relatively recent shake-up of its taxonomy, where the Cretaceous-straddling, globe-spanning monster-Iguanodon genus was carved up into multiple genera spread across time and space (see Darren Naish's Scientific American articles here, here and here for details) did little to revive public interest in one of our longest serving and best-known dinosaurs. Iguanodon seems to be a dinosaurian washed-up Golden Age movie star: once great, now rarely mentioned, and only wheeled for nostalgia.

The gossip magazines would have a field day.

Behind the scenes, however, Iguanodon or, more correctly, 'iguanodonts' are becoming more and more interesting. Far from large, bland and overly-familiar ornithopods, the modern concept of iguanodonts comprises several distinct Lower Cretaceous species with markedly different bauplans which created complex herbivore communities. Their anatomy varied in many aspects other than simply size - even their famous thumb spikes are actually quite disparate - and functionality must have been equally diverse. The very evolution of iguanodonts is also more complex than we thought: rather than forming a clear group of ornithopods, iguanodont taxa seemingly comprise a messy, not-fully-understood bush of species on the ornithopod branch trunk leading to true hadrosaurs (e.g. McDonald 2012a). Thus, there is no truly correct term for a group comprising Iguanodon and its close relatives: 'iguanodont' is used here in a vernacular sense. In short, it seems that iguanodonts have fallen off the popular radar just as they're getting more interesting and worthy of attention

Iguanodonts: the undiscovered country
At the heart of this newfound complexity is the aforementioned reappraisal of iguanodont diversity. It's worth stressing that the charge to slay the waste basket monstergenus Iguanodon, started by Norman and Barrett (2002) and followed by the likes of Paul (2008), Norman (2010), Carpenter and Ishida (2010), Naish and Martill (2008), McDonald et al. (2010), McDonald (2012a, b) and others, was not a case of splitting minor taxonomic hairs. Unlike the differences which separate many fossil animals, most taxa pulled from Iguanodon are characterised by radically different morphology which would be obvious even in life. In Britain alone, the handful of species recognised as various members of Iguanodon may now comprise as many as nine genera (not counting objective synonyms). It's well known that Iguanodon is now monospecific, containing only the giant species I. bernissartensis. In the UK at least, this is principally known from the Wessex Sub-basin of the Wealden Supergroup of the Isle of Wight, although it also occurs in the Weald Sub-basin of Surrey, Sussex and Kent (below). It was joined in both basins by Mantellisaurus, the smaller iguanodont once called Iguanodon atherfieldensis and, in the Wessex, by two other possible taxa: Proplanicoxa galtoni and Dollodon bampingi. All but Proplanicoxa galtoni are known from elsewhere in Europe, which cannot be said for other British iguanodonts Barilium dawsoni*, Hypselospinus fittoni, Sellacoxa pauli and Kukufeldia tilgatensis from the Weald Sub-basin, also of the Wealden Supergroup of Sussex and Surrey. These animals are geologically older than the more familiar Iguanodon and Mantellisaurus and, for now at least, do not seem to overlap stratigraphically. A further genus, Owenodon hoggi, has been named for "Iguanodon" material from the British Purbeck Group. A number of other Asian and North American genera have also been pulled from Iguanodon, but the British record seems unusually diverse and implies that multiple iguanodonts existed in the same basins. Admittedly, exactly how many European iguanodont taxa are valid remains uncertain - there are arguments for it being over-split and overly-conservative - but even a relatively cautious assessment suggests several iguanodont faunas evolved in ancient Britain.

*Fascinating aside: according to Norman (2011a, b) there's a good chance that the original Iguanodon teeth belong to Barilium. There's not much we can do about this now - after years of confusion over what Iguanodon is, the name has been irreversibly transferred to I. bernissartensis. While most agree this was one appropriate cause of action to take - most of us have always thought of this species as the 'classic' Iguanodon - there are lots of niggles and issues with the choice of bernissartensis as the surrogate type species of Iguanodon. The similarity of the original 'I. anglicus' teeth to Barilium is just another hangover from the excessive lumping that Iguanodon experienced in its first 180 years of recognition.

Simplified overview of British iguanodont distribution. The taxa listed here do not include recently named objective synonyms and includes several genera which some authors (e.g. Norman 2011a; McDonald 2012) would happily remove. I. anglicus, the original Iguanodon and nomen dubium, is included for interest only. Silhouettes provide very rough proxies for maximum taxon size to show the possible nature of iguanodont faunas, borrowed from Paul (2008). Hat tip to Bill Wimbledon for some pointers on Wealden chronostratigraphy.

Quite how these contemporary animals did not trip over each others ecological toes remains to be established. Some truth to the 'classic' view of Iguanodon species occurring in different size classes remains, with most newly recognised species equating to large- or medium-size dinosaurian herbivores. What is now very apparent, however, is that size is only one way in which these animals differ. The large iguanodont Barilium, for instance (below), is about the same length as I. bernissartensis (10-12 m) but is much more heavily built, with proportionally heavyset hips, shoulders, limb bones, a chunky anterior tail region and very tall neural spines along much of its back. While it's difficult to call I. bernissartensis a gracile creature, its bones are certainly more svelte than those of Barilium: its limbs are longer, its vertebrae lower, and its limb girdles less stocky. A similar story is echoed in the smaller iguanodonts which lived alongside the giants: Hypselospinus, contemporary of Barilium, was a relatively small (about 6 m long) but stocky species, with chunky limb bones and a thick shoulder girdle. By contrast, other 'small' iguanodonts - such as the 6- 7 m long Mantellisaurus and Dollodon - were rather gracile, with slender limbs and relatively delicate hands. Despite its robust body, Hypselospinus shared a relatively gracile jaw construction with other smaller iguanodonts. With many further differences in their fine anatomy, a clear message can be seen: iguanodonts were not merely resized variants of the same bauplan rolled out over the Lower Cretaceous. Quite how their different anatomies plugged into their palaeoecology and niche differentiation remains to be established, but its possible - maybe probable - that their anatomical differences reflect different foraging strategies, habitat preferences and routine predation responses. Perhaps the geologically younger, slender variants were quicker on their feet than their rotund forebears? Did the more robust species spent more time locomoting quadrupedally? No-one really knows at the moment, but there's clearly a lot of interesting things going on here and a lot of interesting research to be done.

Barilium dawsoni, a large and very robust iguanodont from the Valanginian of Sussex, caked in dried mud. This stunted pollex of this animal, which was probably quadrupedal most of the time, means it'd be hard-pressed to give a thumbs up even if it wanted to. A flock of 'Ashdown maniraptorans', tiny, poorly known theropods no larger than an Eurasian magpie, add scale (see Naish and Sweetman 2011 for details).

It would be remiss of us to not mention that the most famous iguanodont feature - their thumb spikes - are also far from uniform in size or construction. The function of the iguanodont pollex has long proved controversial, but a role in stabbing generic theropods in the neck is a common assumption. This long-held assumption is questioned by the range of morphologies associated with the pollex however. Most of us are familiar with the general construction of the iguanodont pollex thanks to oft-reproduced images of the Iguanodon hand, such as...

Left Iguanodon bernissartensis manus. Image from here.

Here, the pollex is conical and fairly large, but remains detached from the carpal block (iguanodont wrist bones fuse into a single unit with age). Thus, the pollex retains an ability to move somewhat. The pollex of Mantellisaurus is generally similar to that of Iguanodon, except that it is much, much smaller - probably far too small to be used as an effective predator deterrent. By contrast, the pollex of another small iguanodont, Hypselospinus, was proportionally large and robust, being about 40% as long as the forearm. Rather than being truly conical, the pollex of Hypselospinus was laterally compressed and tightly attached to the carpal block so little or no flexion was possible. The thumb of fatso Balirum was actually fused to the block itself, and is of further note for being incredibly short: Barilium would struggle to give a satisfactory 'thumbs up' to anyone. So again, we see evidence of diversity in these unassuming dinosaurs: pollex size, shape, flexion and reinforcement all vary across iguanodont taxa. We may take this as a sign that thumb spike function was also variable across iguanodonts, so there may not be one single explanation for their existence. The tight pollex articulations of some species seemingly make the pollex part of the antebrachial functional unit than the hand, and are strangely reminiscent of the carpometacarpal knobs and spurs of many birds (see - again - a TetZoo series on this topic, starting here). Alas, the function of many bird hand spurs are not well researched, but the general consensus - supported by direct evidence in many cases - is that they're primarily used in combat and aggressive behaviours, much of it intraspecific. In some cases, they may even be used to make noise when slapped against the flanks of their owners. Who knows: perhaps iguanodonts with tightly welded, inflexible thumb spikes used their pollices in a similar way. But what of species with flexible thumb spikes? Could they be used as weapons too? If so, how come the large pollex of Iguanodon was not fused to the carpus when the large thumb of Hypselospinus is? Did that make it a less effective weapon? And what was Mantellisaurus using that piddling little thumb spike for, if anything? Questions, questions, questions...

The bit where I stop writing
In sum, while it would be silly to say that iguanodont science is undergoing anything like a revolution or renaissance, there's certainly a lot of tinkering going on and the results are exciting whatever your specific taste in palaeontology - taxonomic, functional, or palaeoecological. Granted, the outcome of these ongoing studies are not going to make newspaper headlines, but if you're interested in dinosaur palaeobiology - and you are if you've read this far - then this should be very cool, interesting stuff. If the apparent decline in public interest for iguanodonts is because many of us consider them overly-familiar, then we need to think about changing that attitude. Far from being 'done to death', after many decades of fairly static interpretation, iguanodont science is becoming more interesting than ever.

For an easy to access, relatively up to date and inexpensive look at a bunch of iguanodonts, you could do a lot worse than checking out Dave Norman's chapter on ornithopods in English Wealden Fossils (Norman, 2011b). Further brief musings on the decline of a dinosaur celebrity are provided in this post on Stegosaurus.

References

• Carpenter, K., & Ishida, Y. (2010). Early and “Middle” Cretaceous iguanodonts in time and space. Journal of Iberian Geology, 36(2), 145-164.
• Paul, G. S. (2008). A revised taxonomy of the iguanodont dinosaur genera and species. Cretaceous Research, 29(2), 192-216.
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